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ISSN 0568-5435 ÐÎÑÑÈÉÑÊÀß ÀÊÀÄÅÌÈß ÍÀÓÊ ÁÎÒÀÍÈ×ÅÑÊÈÉ ÈÍÑÒÈÒÓÒ èì. Â. Ë. ÊÎÌÀÐÎÂÀ ACADEMIA SCIENTIARUM ROSSICA INSTITUTUM BOTANICUM NOMINE V. L. KOMAROVII ÍÎÂÎÑÒÈ ÑÈÑÒÅÌÀÒÈÊÈ ÍÈÇØÈÕ ÐÀÑÒÅÍÈÉ ÒÎÌ 43 NOVITATES SYSTEMATICAE PLANTARUM NON VASCULARIUM TOMUS XLIII Òîâàðèùåñòâî íàó÷íûõ èçäàíèé KMK Ñàíêò-Ïåòåðáóðã Ìîñêâà v 2009 À. Ã. Øèðÿåâ A. G. Shiryaev ÊËÀÂÀÐÈÎÈÄÍÛÅ ÃÐÈÁÛ ÒÓÍÄÐÎÂÎÉ È ËÅÑÎÒÓÍÄÐÎÂÎÉ ÇÎÍ ÊÎËÜÑÊÎÃÎ ÏÎËÓÎÑÒÐÎÂÀ (ÌÓÐÌÀÍÑÊÀß ÎÁËÀÑÒÜ) CLAVARIOID FUNGI OF THE TUNDRA AND FOREST-TUNDRA ZONES OF KOLA PENINSULA (MURMANSK REGION) Èíñòèòóò ýêîëîãèè ðàñòåíèé è æèâîòíûõ ÓðÎ ÐÀÍ Ëàáîðàòîðèÿ ôèòîìîíèòîðèíãà è îõðàíû ðàñòèòåëüíîãî ìèðà 620144, Åêàòåðèíáóðã, óë. 8 Ìàðòà, ä. 202 [email protected] Ïÿòüäåñÿò øåñòü âèäîâ êëàâàðèîèäíûõ ãðèáîâ èç 14 ðîäîâ îòìå÷åíû â âû- ñîêîøèðîòíûõ ðàéîíàõ Êîëüñêîãî ïîëóîñòðîâà, èç íèõ 55 âûÿâëåíû â ëåñîòóí- äðå è 30 â þæíûõ òóíäðàõ. Äâà âèäà (Mucronella flava è Ramaria testaceoflava) âïåðâûå îòìå÷åíû â ëåñîòóíäðîâîé çîíå Ðîññèè. Íàèáîëüøåå âèäîâîå áîãàòñòâî îòìå÷åíî äëÿ ðîäà Typhula (19 âèäîâ), à òðè êðóïíåéøèõ ðîäà Typhula, Ramaria è Clavaria âêëþ÷àþò â ñóììå 53.6% âñåõ âûÿâëåííûõ âèäîâ. Êîýôôèöèåíò âè- äîâîé íàñûùåííîñòè ðîäà â ëåñîòóíäðå ðàâåí 4.0, à â þæíûõ òóíäðàõ 3.7. Óðîâåíü âèäîâîãî áîãàòñòâà, ðàçíîîáðàçèÿ (èíäåêñ Øåííîíà) è äîìèíèðîâàíèÿ (èíäåêñ Ñèìïñîíà) ñõîäíû ñ òàêîâûìè äëÿ äðóãèõ âûñîêîøèðîòíûõ ðåãèîíîâ Åâðîïû, îäíàêî íåñêîëüêî îòëè÷àþòñÿ îò àíàëîãè÷íûõ ïîêàçàòåëåé äëÿ Ïîëÿð- íîãî Óðàëà è ßìàëà. Íàèáîëåå îáèëüíûìè âèäàìè ÿâëÿþòñÿ ñàïðîòðîôû, ðàç- âèâàþùèåñÿ íà òðàâÿíèñòîì îïàäå Typhula variabilis, T. lutescens è T. sclero- tioides. Ïî ñðàâíåíèþ ñ àíàëîãè÷íûìè êîìïëåêñàìè Ïîëÿðíîãî Óðàëà è ßìàëà íà Êîëüñêîì ïîëóîñòðîâå çíà÷èòåëüíà ðîëü ãóìóñîâûõ ñàïðîòðîôîâ (Clavulina cinerea, Clavaria argillacea, C. falcata). Êñèëîòðîôíûå âèäû îòìå÷åíû ëèøü â ëåñîòóíäðå. Êëþ÷åâûå ñëîâà: Åâðàçèÿ, êëàâàðèîèäíûå ãðèáû, Êîëüñêèé ïîëóîñòðîâ, ëå- ñîòóíäðà, ñóáàðêòèêà, òóíäðà. Fifty six species of clavarioid fungi from 14 genera are noted in the high-latitude areas of Kola Peninsula, among them 55 species being noted in forest-tundra and 30 in southern tundra. Two species (Mucronella flava and Ramaria testaceoflava) are recorded in the Russian forest-tundra zone for the first time. Like in all high-altitude areas of Eurasia, the greatest species richness is distinguished for the genus Typhula (19 species), and 3 largest genera Typhula, Ramaria and Clavaria make in total 53.6% of all revealed taxa. The species/genus ratio in forest-tundra is 4.0, and in southern tundra 3.7. The levels of species richness, diversity (Shannons index) and domination (Simpsons index) are similar to those for other European high-latitude areas, however somewhat differ from similar parameters for Polar Urals and Yamal Peninsula. Saprotrophic species on grass litter Typhula variabilis, T. lutescens and T. sclerotioides are the most abundant. In comparison with other high-latitude complexes, the role of 134 soil-inhabiting saprotrophs (Clavulina cinerea, Clavaria argillacea, C. falcata) is significant on Kola Peninsula. Wood-decaying species are noted only in forest-tundra. Keywords: clavarioid fungi, forest-tundra, Eurasia, Kola Peninsula, Subarctic, tundra. Âûñîêîøèðîòíûé êîìïëåêñ êëàâàðèîèäíûõ ãðèáîâ ðîññèéñêîé Àðêòèêè èçó÷åí ñëàáî (Êàðàòûãèí è äð., 1999; Shiryaev, 2006), â òîì ÷èñëå è â Ìóðìàíñêîé îáëàñòè, ãäå ïî îïóáëèêîâàííûì äàííûì èç- âåñòíî 11 âèäîâ êëàâàðèîèäíûõ ãðèáîâ (Karsten, 1882; Ïàðìàñòî, 1965; Øóáèí, Êðóòîâ, 1979; Êîòêîâà, 2007; Shiryaev, Kotiranta, 2007). Îäíàêî ïîäàâëÿþùàÿ ÷àñòü âûÿâëåííûõ ðàíåå ãðèáîâ ýòîé ãðóïïû ñîáðàíà â òàåæíûõ ðàéîíàõ îáëàñòè, è ëèøü ÷åòûðå èç íèõ áûëè âñòðå÷åíû â òóíäðîâîé è ëåñîòóíäðîâîé çîíàõ (Shiryaev, Kotiranta, 2007). Ñîãëàñíî çîíàëüíîìó ðàéîíèðîâàíèþ Àðêòèêè (Êàìåëèí, 2004), êðàéíèé ñåâåð Êîëüñêîãî ïîëóîñòðîâà, ïîëíîñòüþ ëåæàùåãî çà Ïî- ëÿðíûì êðóãîì, îòíîñèòñÿ ê ïîäçîíå þæíûõ ãèïîàðêòè÷åñêèõ òóíäð è ëåñîòóíäðîâîé çîíå. Îñîáåííîñòè êëèìàòà ýòîé òåððèòîðèè îáóñ- ëîâëåíû âëèÿíèåì òåïëîãî òå÷åíèÿ Ãîëüôñòðèì, áëàãîäàðÿ êîòîðîìó äàæå â ñóðîâûå çèìû þãî-çàïàäíàÿ ÷àñòü ïðèëåæàùåãî Áàðåíöåâà ìîðÿ íå çàìåðçàåò, ïîýòîìó âîïðîñ îòíåñåíèÿ ýòîé òåððèòîðèè ê àð- êòè÷åñêèì ðàéîíàì äî íàñòîÿùåãî âðåìåíè ÿâëÿåòñÿ äèñêóññèîííûì (×èíåíêî, 2008). Êëèìàò ñåâåðíîé ÷àñòè Êîëüñêîãî ï-îâà îòëè÷àåòñÿ âûñîêîé âëàæ- íîñòüþ, ÷àñòîé îáëà÷íîñòüþ, ãóñòûìè òóìàíàìè è îòòåïåëÿìè çèìîé. Ñðåäíåãîäîâàÿ òåìïåðàòóðà âîçäóõà â þæíûõ òóíäðàõ êîëåáëåòñÿ â ðàéîíå 2 °Ñ, ñ íàèáîëåå íèçêîé òåìïåðàòóðîé â ôåâðàëå (îò 5 äî 9 °Ñ ), à íàèáîëåå âûñîêîé â èþëå (9 °Ñ). Áåçìîðîçíûé ïåðèîä ïðîäîëæàåòñÿ 100120 äíåé. Ñðåäíÿÿ ãîäîâàÿ âëàæíîñòü âîçäóõà êî- ëåáëåòñÿ â ïðåäåëàõ 7680%, à ãîäîâîå êîëè÷åñòâî îñàäêîâ îò 500 äî 700 ìì, ïðåèìóùåñòâåííî âûïàäàþùèõ â òåïëîå âðåìÿ ãîäà â èþíå è àâãóñòå.  ëåñîòóíäðå ñðåäíåãîäîâàÿ òåìïåðàòóðà ñîñòàâëÿ- åò îêîëî 2.5 °Ñ ñ íàèáîëåå íèçêîé òåìïåðàòóðîé â ÿíâàðå (îò 8 äî 11 °Ñ), è ñ íàèáîëåå âûñîêîé â èþëå (11 °Ñ). Ïîëÿðíàÿ çèìà ïðî- äîëæàåòñÿ 56 ìåñÿöåâ, íî îíà îòíîñèòåëüíî òåïëàÿ ïî ñðàâíåíèþ ñ çèìîé â äðóãèõ îáëàñòÿõ, ðàñïîëîæåííûõ â òåõ æå øèðîòàõ. Þæíûå òóíäðû çàíèìàþò íå áîëåå 20% ïëîùàäè Ìóðìàíñêîé îáë., îãèáàÿ Êîëüñêèé ï-îâ ñ ñåâåðî-çàïàäà íà þãî-âîñòîê ïîëîñîé â 2030 êì øèðèíîé. Çäåñü ïðåîáëàäàåò êóñòàðíèêîâàÿ òóíäðà (Betula nana, Salix lapponicum, S. glauca è äð.) ëèøàéíèêîâîãî è ìîõîâîãî 135 òèïà. Áëèæå ê ìîðþ ñðåäè êóñòàðíèêîâ âñòðå÷àåòñÿ ìíîãî Rubus chamaemorus, Vaccinium myrtilis, V. vitis-idaea è äð. Áîëüøèå ïðî- ñòðàíñòâà íà äðåíèðîâàííûõ ó÷àñòêàõ çàíÿòû øèêøåâîé òóíäðîé (Empetrum nigrum) ãäå âñòðå÷àþòñÿ Chamaepericlymenum suecicum, Arctostaphylos alpina, Calluna vulgaris, Loiseleuria procumbens, Dryas punctata, Salix herbaceus, Saussurea alpina, Chamaenerion angusti- folium. Ôðàãìåíòû âûñîêîòðàâüÿ ïðåäñòàâëåíû Filipendula ulmaria, Heracleum sp., Anthriscus sylvestris, Valeriana officinalis. Êðîìå òîãî, ìíîãî ñôàãíîâûõ áîëîò è ðó÷üåâ, ïî îêðàèíàì êîòîðûõ ìíîãî÷èñëåí- íû çàðîñëè èâíÿêà. Ëåñîòóíäðà îáðàçóåò ïîëîñó äî 100 êì øèðèíîé. Ïî÷òè âåçäå îíà ïðåäñòàâëåíà áåðåçîâûì ðåäêîëåñüåì, èíîãäà ñ åëüþ (Picea abies) èëè ñîñíîé (Pinus sylvestris), ïðåèìóùåñòâåííî ëèøàé- íèêîâîãî òèïà. Øèðîêî ïðåäñòàâëåíî ðàçíîòðàâüå: Carex spp., Deschamsia flexuosa, Equisetum pratense, Tofieldia palustris, Silene acaulis, Rumex sp., Stellaria sp., Athyrium sp. è âûñîêîòðàâüå: Filipendula ulmaria, Heracleum sp., Anthriscus sylvestris. Èíòðàçîíàëü- íàÿ ðàñòèòåëüíîñòü â îñíîâíîì ïðåäñòàâëåíà èâíÿêàìè, èíîãäà ïåðå- ìåæàþùèìèñÿ ðÿìîâûìè çàðîñëÿìè ñîñíû íà òîðôÿíûõ áîëîòàõ. Äëÿ ãîðíûõ ìàññèâîâ Õèáèí õàðàêòåðíû ñóáàëüïèéñêèå òóíäðû íà âåðøèíàõ. Ïîëåâûå ðàáîòû ïðîâîäèëèñü â ñåíòÿáðå 1998 ã. â ÷åòûðåõ îñíîâ- íûõ òî÷êàõ (ðèñ. 1), äâå èç êîòîðûõ ðàñïîëîæåíû â çîíå þæíûõ êó- ñòàðíè÷êîâûõ òóíäð: 1 â îêðåñòíîñòè ïîñ. Ëèèíàõàìàðè (69°39′ ñ. ø., 31°23′ â. ä.) è 2 â îêðåñòíîñòè ïîñ. Ïîëÿðíûé (69°11′ ñ. ø., 33°29′ â. ä.), à äâå â ëåñîòóíäðå: 3 â îêðåñòíîñòè îç. Ñàíòàÿð- âè (69°30′ ñ. ø., 31°20′ â. ä.) è 4 â îêðåñòíîñòè îç. Íüÿë (69°08′ ñ. ø., 32°15′ â. ä.).  ñõîäíûå ñðîêè áûëî îáñëåäîâàíî ìàêñèìàëüíî ïîëíîå ÷èñëî ïîòåíöèàëüíûõ òèïîâ ñóáñòðàòîâ (ìåñòîîáèòàíèé) äëÿ êàæäîãî ïðèðîäíî-çîíàëüíîãî êîìïëåêñà. Ïîä îáðàçöîì (åäèíèöåé ó÷åòà) ïîíèìàåòñÿ ãðóïïà (êîëîíèÿ, ñêîïëåíèå) ïëîäîâûõ òåë âèäà, îòñòîÿùàÿ îò äðóãîé àíàëîãè÷íîé ãðóïïû ýòîãî âèäà íå ìåíåå ÷åì íà 15 ì. Îáíàðóæåíèå áàçèäèîì âèäà â ãðóïïå ðàññìàòðèâàåòñÿ êàê îäíà åäèíèöà ó÷åòà íåçàâèñèìî îò êîëè÷åñòâà áàçèäèîì è ðàçìåðà ãðóï- ïû. Äëÿ îöåíêè îáèëèÿ ðàññ÷èòàí ïîêàçàòåëü âñòðå÷àåìîñòè (Íîâî- æèëîâ, 2005): P = n / N, i i ãäå ãäå n ÷èñëî ðåãèñòðàöèé (åäèíèö ó÷åòà) êàæäîãî âèäà, N i îáùåå ÷èñëî åäèíèö ó÷åòà âñåõ âèäîâ â àíàëèçèðóåìîì íàáîðå äàí- íûõ. 136 Ðèñ. 1. Ðàéîí èññëåäîâàíèÿ è ïóíêòû ñáîðà ìàòåðèàëà â þæíûõ òóíäðàõ è ëåñîòóíäðå Êîëüñêîãî ïîëóîñòðîâà. 1 îêð. ïîñ. Ëèèíàõàìàðè; 2 îêð. ïîñ. Ïîëÿðíûé, 3 îêð. îç. Ñàíòàÿðâè, 4 îêð. îç. Íüÿë. Âèäîâîå ðàçíîîáðàçèå âû÷èñëåíî ñ ïîìîùüþ èíäåêñà Øåííîíà: H′ = Σ P log P , i i ãäå P âñòðå÷àåìîñòü i-ãî âèäà (Magguran, 2004). Ïðè ñðàâíåíèè i ðàçíîîáðàçèÿ ïîäñ÷èòàí êðèòåðèé Ñòüþäåíòà (t). Äëÿ âû÷èñëåíèÿ äîìèíèðîâàíèÿ èñïîëüçîâàí èíäåêñ Ñèìïñîíà: D = Σ P 2. i Äëÿ ñðàâíåíèÿ âèäîâîãî ñîñòàâà ðàçëè÷íûõ ñîîáùåñòâ èñïîëüçî- âàí êîýôôèöèåíò Ѹðåíñåíà ×åêàíîâñêîãî: C = 2C / (A+B), s ãäå À îáùåå êîëè÷åñòâî âèäîâ â ïåðâîì ñîîáùåñòâå,  îáùåå êîëè÷åñòâî âèäîâ âî âòîðîì ñîîáùåñòâå, Ñ êîëè÷åñòâî âèäîâ, îá- ùèõ äëÿ ñðàâíèâàåìûõ ñîîáùåñòâ. Äåíäðîãðàììà ïîñòðîåíà ñ èñïîëüçîâàíèåì ïðîãðàììû Statistica 6.0, ãäå èñïîëüçîâàí ìåòîä êëàñòåðèçàöèè Âàðäà, à â êà÷åñòâå ìåðû ðàçëè÷èé âûñòóïàåò êîýôôèöèåíò Ѹðåíñåíà ×åêàíîâñêîãî. 137 Òàáëèöà 1 Âèäîâîå ðàçíîîáðàçèå è áîãàòñòâî áèîòû êëàâàðèîèäíûõ ãðèáîâ ëåñîòóíäð è þæíûõ òóíäð Êîëüñêîãî ïîëóîñòðîâà Ëåñîòóíäðà Þæíûå òóíäðû ×èñëî îáðàçöîâ 291 149 ×èñëî âèäîâ 55 30 ×èñëî ðîäîâ 13 8 Â/Ð 4.0 3.7 H' 3.88 3.34 D 0.021 0.039 Ïðèìå÷àíèå. Â/Ð êîýôôèöèåíò âèäîâîé íàñûùåííîñòè ðîäà; H′ èíäåêñ ðàç- íîîáðàçèÿ Øåííîíà; D èíäåêñ äîìèíèðîâàíèÿ Ñèìïñîíà.  ðåçóëüòàòå èññëåäîâàíèé âûÿâëåíî 440 îáðàçöîâ (åäèíèö ó÷å- òà), âêëþ÷àþùèõ 243 ïîëåâûõ îáðàçöà è 197 çàïèñåé â ïîëåâîé äíåâ- íèê, â òîì ÷èñëå â ëåñîòóíäðå 291 åäèíèöà ó÷åòà, â þæíûõ òóí- äðàõ 149 (òàáë. 1). Ñîáðàííûé ìàòåðèàë õðàíèòñÿ â êîëëåêöèè àâòîðà, íàõîäÿùåéñÿ â ìèêîëîãè÷åñêîì ãåðáàðèè Èíñòèòóòà ýêîëîãèè ðàñòåíèé è æèâîòíûõ ÓðÎ ÐÀÍ â Åêàòåðèíáóðãå (SVER). Íàçâàíèÿ êëàâàðèîèäíûõ ãðèáîâ è ñîêðàùåíèÿ ôàìèëèé àâòîðîâ äàíû â ñîîòâåòñòâèè ñ áàçîé äàííûõ Index Fungorum (http:// www.indexfungorum.org/Names/Names.asp) (íà 15.04.2009), âèäû ðàñ- ïîëîæåíû â àëôàâèòíîì ïîðÿäêå. Íîâûå äëÿ Ìóðìàíñêîé îáë. âèäû îòìå÷åíû çâåçäî÷êîé (*). Äëÿ êàæäîãî âèäà ïðèâîäÿòñÿ: íàçâàíèå,